Consumption of a focal plant by herbivores depends, not only on the physical and chemical characteristics of that plant, but also on the characteristics of the neighbouring vegetation. Consumption of focal plants has been related to their own characteristics and to the quality of the neighbouring vegetation, but the two have not been combined to examine the relative importance of focal plant and neighbouring vegetation characteristics. We conducted a series of feeding trials to examine the relative importance of focal plant and various characteristics of neighbouring vegetation to browsing of a focal plant within vegetation patches. We planted Eucalyptus nitens seedlings of high and low nutrient status amongst vegetation patches differing in palatability, abundance and height. Generalist mammalian herbivores, red‐bellied pademelons (Thylogale billardierii), were allowed to feed in each of these patches one at a time, and seedling consumption was recorded. Results were considered in light of the attractant‐decoy and apparency hypotheses, which focus on the outcome to plants, and in terms of foraging theory, which is process‐focussed. Seedling and vegetation characteristics were both important. Seedlings of high nutrient status were preferred over those of low nutrient status. The relative quality, abundance and height of neighbouring vegetation all influenced browsing of a focal plant. Seedlings were more vulnerable amongst vegetation that was of low palatability, of low abundance, or was short. Seedling and vegetation effects were additive in two of three trials. Results were consistent with both the attractant‐decoy and apparency hypotheses, and could be explained in terms of maximising foraging efficiency. They demonstrate the need to consider characteristics of both the focal plant and its neighbouring vegetation when predicting the vulnerability of the former to browsing by generalist herbivores.
Summary 1.Foraging decisions by herbivores depend on variation in food types, the scale(s) at which this variation occurs and the opportunity and capacity for herbivores to respond to such variation. These decisions affect not only the herbivores themselves, but also the vulnerability of individual plants to being eaten. Associational plant refuges, in which neighbouring plants alter focal plant vulnerability, are an emergent property of foraging decisions. 2. Using the red-bellied pademelon (Thylogale billardierii) as a model generalist mammalian herbivore, we investigated the spatial scale(s) at which animals made foraging decisions and the resultant effect on focal plant vulnerability. In a replicated design, we varied vegetation at the individual plant scale, generating intraspecific differences in Eucalyptus nitens seedlings by altering their nutrient status (high, low). We varied vegetation at the patch scale, in which seedlings were planted, using high-(grass) and low-(herbicided) quality patches. Animals were allowed to choose where they fed and what they ate. Animal behaviour was recorded and intake of seedlings measured. 3. We found that animals made foraging decisions first at the patch scale then at the scale of individual plants; both patch and focal seedling characteristics influenced browsing. Pademelons spent most of their time in high-quality patches, and seedlings were consequently more vulnerable there than in low-quality patches. Pademelons also ate more foliage from high-than from low-nutrient status seedlings. This behaviour concentrated resources, increasing foraging efficiency and making focal plants more vulnerable to browsing. 4. The opportunity and capacity to choose at both plant and patch scales resulted in a pattern of focal plant vulnerability consistent with the repellent-plant hypothesis. This contrasts with our previous study, in which animals were only provided with choice at the plant level and plant vulnerability followed the attractant-decoy hypothesis. These combined results demonstrate that the influence of neighbouring vegetation on consumption of a focal plant depends on the spatial scale of selection and on opportunities (and capacity) for herbivores to choose.
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